Pleistocene diversification of the Pomacentrus coelestis species complex (Pisces: Pomacentridae): historical biogeography and species boundaries

Pleistocene eustatic changes in sea level are often invoked to explain genetic divergence among marine organisms. However, molecular phylogenies have revealed relatively few examples of speciation events dating to the Pleistocene. We present a species-level hypothesis of the timing of evolution for the Pomacentrus coelestis species complex (Pomacentridae), based on the nuclear S7 intron and the mitochondrial Cytb gene, and reconstruct ancestral range distributions across the timetree. Ancestral range reconstruction suggests the complex originated in the Coral Triangle and East Indian Ocean, with subsequent range expansion outward from this region. We suggest that land barriers to dispersal (e.g., Indo-Pacific barrier) may be responsible for the divergence between Indian (P. alleni, P. similis, P. caeruleopunctatus, and P. caeruleus) and Pacific (P. micronesicus, P. auriventris, and P. coelestis) species groups, and subsequent isolation by Pleistocene sea-level fluctuations in certain areas of the Coral Triangle (glacial refugia) may play an important role in the diversification of this species complex. Additionally, our analyses show cryptic lineages within P. micronesicus and highlight the need for comprehensive sampling within and among species to reveal recent speciation events.     

Dynamics of larval fish assemblages over a shallow coral reef in the Florida Keys

Few time series collections have been made of the larval ichthyofauna in waters directly above shallow coral reefs. As a result, relatively little is known regarding the composition and temporal dynamics of larval fish assemblages in shallow-reef waters, particularly those near a major western boundary current. We conducted a series of nightly net tows from a small boat over a shallow reef (Pickles Reef) along the upper Florida Keys during four new moon and three third-quarter moon periods in July (two new moons), August, and September 2000. Replicate tows were made after sunset at 0–1 m and at 4–5 m depth to measure the nightly progression in community composition, differences in depth of occurrence, and abundance and diversity with lunar phase. A total of 66 families was collected over the 3-month period, with a mean (±SE) nightly density of 23.7±2.1 larvae per 100 m ³ and diversity of 24.2±0.9 taxa per tow. A total of 28.8% of the catch was composed of small, schooling fishes in the families Atherinidae, Clupeidae, and Engraulidae. Of the remaining catch, the top ten most abundant families included reef fishes as well as mangrove and oceanic taxa (in descending order): Scaridae, Blennioidei (suborder), Gobiidae, Paralichthyidae, Lutjanidae, Haemulidae, Labridae, Gerreidae (mangrove), Balistidae, and Scombridae (oceanic). These near-reef larval fish assemblages differed substantially from those collected during previous offshore collections. Taxa such as the Haemulidae were collected at a range of sizes and may remain nearshore throughout their larval period. Overall, the abundance and diversity of taxa did not differ with depth (although within-night vertical migration was evident) or with lunar phase. Temporal patterns of abundance of larval fish families clustered into distinct groups that in several cases paralleled family life-history patterns. In late July, a sharp shift in larval assemblages signaled the replacement of oceanic water with inner shelf/bay water. In general, the suite and relative abundance of taxa collected each night differed from those collected on other nights, and assemblages reflected distinct nightly events as opposed to constant or cyclical patterns. Proximity to the Florida Current likely contributes to the dynamic nature of these near-reef larval assemblages. Our results emphasize the uniqueness of near-reef larval fish assemblages and point to the need for further examination of the biophysical relationships generating event-related temporal patterns in these assemblages.     

Planktonic larval duration of one hundred species of Pacific and Atlantic damselfishes (Pomacentridae)

The plankton larval duration for 100 species of Pacific and Atlantic damselfishes was estimated from daily growth increments on the otolith of juvenile fish collected at various localities between July 1987 and September 1988. For newly-settled fishes, larval duration was determined by counting the entire number of increments present on the otolith, while for older juveniles estimates were made by counting the number of increments between the center of the otolith and a mark corresponding to settlement. We document the development of otolith formation during the period when eggs are incubated on the reef and show that daily increments are only accreted after larvae hatch and enter the planktonic phase. The planktonic larval duration for damselfish is shorter and less variable, both between and within species, compared to other groups of reef fishes such as wrasses and surgeonfishes. Larval duration ranged from 12 to 39 d. Average duration between species ranged from 13.1 to 35.2 d. The time spent in the plankton was not significantly correlated with geographic distribution when evaluated among species, however, genera with confined regional distribution have a shorter mean larval life than do widely distributed genera. Size at settlement was positively correlated with time spent in the plankton among species, but a significant correlation between these variables was only evident within one of ten species. The low variance in planktonic larval duration within species indicates that most damselfish are unable to delay metamorphosis following competency. This inability to postpone settlement limits the potential for dispersal, especially when dispersal time between suitable habitats is greater than about 30 d.     

Scale-dependent changes in the importance of larval supply and habitat to abundance of a reef fish

While there is great interest in the degree to which local interactions "scale-up" to predict regional patterns of abundance, few studies in marine systems have simultaneously examined patterns of abundance at both the large scale (tens of kilometers) typical of larval movement and the small scale (meters) typical of post-settlement interactions. We addressed this gap by monitoring larval supply, adult survivorship, and giant kelp (Macrocystis pyrifera, a primary habitat-forming species) abundance for 13 populations of kelp bass (Paralabrax clathratus) spread over approximately 200 km in the Santa Barbara Channel, California, USA. At the small, within-site scale, both recruitment and adult survivorship of kelp bass were density-dependent and positively related to kelp abundance. At the larger, among-site scale, the spatial pattern of adult kelp bass abundance was predicted well by the pattern of kelp bass larval supply, but there was a consistent negative spatial relationship between kelp abundance and kelp bass larval supply despite the positive effects of kelp on kelp bass at the smaller spatial scale. This large-scale negative relationship was likely a product of a channel-wide spatial mismatch between oceanographic conditions that favor kelp survival and those that concentrate and distribute fish larvae. These results generally support the recruit-adult hypothesis: kelp bass populations are limited by recruitment at low recruit densities but by density-dependent competition for food resources and/or predator refuges at high recruit densities. At the same time, spatial variation in kelp abundance produced substantial spatiotemporal heterogeneity in kelp bass demographics, which argues for a multispecies, metacommunity approach to predicting kelp bass dynamics.     

Auditory and olfactory abilities of pre-settlement larvae and post-settlement juveniles of a coral reef damselfish (Pisces: Pomacentridae)

The propagules of most species of reef fish are advected from the reef, necessitating a return to reef habitats at the end of the pelagic stage. There is increasing evidence of active attraction to the reef but the sensory abilities of reef fish larvae have not been characterized well enough to fully identify cues. The electrophysiological methods of auditory brainstem response (ABR) and electroolfactogram (EOG) were used to investigate auditory and olfactory abilities of pre- and post-settlement stages of a damselfish, Pomacentrus nagasakiensis (Pisces, Pomacentridae). Audiograms of the two ontogenetic stages were similar. Pre-settlement larvae heard as well as their post-settlement counterparts at all but two of the tested frequencies between 100 Hz and 2,000 Hz. At 100 and 600 Hz, pre-settlement larvae had ABR thresholds 8 dB higher than those of post-settlement juveniles. Both stages were able to detect locally recorded reef sounds. Similarly, no difference in olfactory ability was found between the two ontogenetic stages. Both stages showed olfactory responses to conspecifics as well as L-alanine. Therefore, the auditory and olfactory senses have similar capabilities in both ontogenetic stages. Settlement stage larvae of P. nagasakiensis can hear and smell reef cues but it is unclear as to what extent larvae use these sounds or smells, or both, as cues for locating settlement sites. An erratum to this article can be found at     

Feeding macroecology of territorial damselfishes (Perciformes: Pomacentridae)

The present study provides the first analysis of the feeding macroecology of territorial damselfishes (Perciformes: Pomacentridae), a circumtropical family whose feeding and behavioral activities are important in structuring tropical and subtropical reef benthic communities. The analyses were conducted from data collected by the authors and from the literature. A strong positive correlation was observed between bite rates and sea surface temperature (SST) for the genus Stegastes. A negative correlation was found between bite rates and mean body size for the genera Stegastes and Pomacentrus, but this relationship was not significant when all territorial pomacentrids were analyzed together. A negative correlation between body size and SST was observed for the whole group and for the genera Stegastes, and Pomacentrus. No relationship was found between territory size and feeding rates. Principal Components Analysis showed that differences in feeding rates accounted for most of the variability in the data. It also suggested that body size may be important in characterizing the different genera. In general, tropical species are smaller and have higher bite rates than subtropical ones. This study extended the validity of Bergmann’s rule, which states that larger species or larger individuals within species occur towards higher latitudes and/or lower temperatures, for an important group of reef fishes. The identification of large-scale, robust ecological patterns in the feeding ecology of pomacentrid fishes may establish a foundation for predicting large-scale changes in reef fish assemblages with expected future changes in global SST.     

Wave exposure,swimming performance,and the structure of tropical and temperate reef fish assemblages

We examined the relationship between swimming performance, wave exposure, and the distribution patterns of labrids on temperate rocky reefs, in comparison with previous functional analyses of a tropical assemblage. Visual censuses of the distribution and abundance of labrids across two major gradients of wave exposure (depth and aspect to prevailing winds) were made at two offshore islands near Port Stephens, New South Wales, Australia. Distinct shifts in species composition and abundance were evident between high and low wave exposure habitats on temperate rocky reefs, particularly between deep and shallow habitats on exposed reef fronts. The swimming performances of temperate labrids were assessed through examination of pectoral fin shape (aspect ratio) and in situ swimming speeds. A diversity of pectoral fin morphologies was exhibited within this temperate assemblage, ranging from rounded to tapered fins (aspect ratios of 0.52 and 1.43, respectively). Fin shape was strongly correlated (Pearsons correlation 0.884, P<0.001) with swimming speed (ranging from 1.05 and 3.06 body lengths s^–1), in a relationship comparable to that observed in tropical labrids. Inter-specific differences in swimming ability provided some explanation for differences in the distribution and abundance of temperate labrids in relation to wave exposure. However, our findings suggest that although coral reef labrids appear to predominantly use high aspect-ratio fins to successfully occupy wave-exposed habitats, temperate labrids appear to be using an enhanced swimming ability through increased body size.Communicated by G.F. Humphrey, Sydney     

Variability in nursery function of tropical seagrass beds during fish ontogeny: timing of ontogenetic habitat shift

Seagrass beds are often considered to be important nurseries for coral reef fish, yet the effectiveness of these nursery functions (refuge and food availability) at different juvenile stages is poorly understood. To understand how the demands of juvenile fish on seagrass nursery functions determines the timing of ontogenetic habitat shifts from seagrass beds to coral reefs, we conducted visual transect survey and field tethering and caging experiments on three different sizes of the coral reef fish Pacific yellowtail emperor (Lethrinus atkinsoni) during its juvenile tenure in seagrass beds at Ishigaki Island, southern Japan. The study showed that although the number of individual L. atkinsoni juveniles decreased by >90 % during their stay in the seagrass nursery, the shelter and/or food availability functions of the nursery, at least for a juvenile size of approximately 5 cm total length (TL), provided the best survival and growth option. The timing of ontogenetic migration to coral reefs of larger fish (>8 cm TL) was attributed to foraging efficiency for larger food items in different habitats. Overall, the function of the seagrass bed nursery changed with juvenile body size, with marginally higher survival and significantly greater growth rates during early juvenile stages in seagrass beds compared to coral reefs. This would contribute to the enhancement in the number of individuals eventually recruited to adult populations.     

Reproduction and larval development of Marenzelleria viridis (Polychaeta: Spionidae)

The pelagic larvae of the polychaete Marenzelleria viridis (Verrill, 1873) are newcomers among the meroplankton of the Baltic Sea, where they sometimes achieve an abundance of up to 21x10⁶ ind m^-3 near the coast, especially in late autumn and even in winter. Benthos samples were collected in the Darss-Zingst bodden chain from April 1992 to 1993 and used to ascertain the reproductive stages of individuals. Development from fertilized egg to benthic juvenile is described on the basis of field material. Details of the morphology of various pelagic stages and the young benthic worm are presented as drawings and scanning electron microscope photographs. In 1992, development of the gametes started in mid-May. The individuals reached maturity in late September after about 20 wk. The first planktonic stage was the fertilized egg. The larvae have initially one and later two pairs of black eyes and, in the 2nd segment, a ciliated pit. Gastrotrochs are present on the third and thereafter every alternate segment. Neuropodial setae develop once the 7-setiger stage is reached. Palps appear at the 10-setiger stage, and neuropodial bidentate hooks from the 10th to 11th setiger on. Metamorphosis into the juvenile benthic stage takes place at the earliest when the 15-setiger stage has been reached. The reproductive seasons of various populations of M. viridis and the developmental patterns of their larvae are discussed, and differences are compared with relevant findings for the spionids.     

Connectivity between marine reserves and exploited areas in the philopatric reef fish Chrysoblephus laticeps (Teleostei: Sparidae)

‘No-take’ marine protected areas (MPAs) are successful in protecting populations of many exploited fish species, but it is often unclear whether networks of MPAs are adequately spaced to ensure connectivity among reserves, and whether there is spillover into adjacent exploited areas. Such issues are particularly important in species with low dispersal potential, many of which exist as genetically distinct regional stocks. The roman, Chrysoblephus laticeps, is an overexploited, commercially important sparid endemic to South Africa. Post-recruits display resident behavior and occupy small home ranges, making C. laticeps a suitable model species to study genetic structure in marine teleosts with potentially low dispersal ability. We used multilocus data from two types of highly variable genetic markers (mitochondrial DNA control region and seven microsatellite markers) to clarify patterns of genetic connectivity and population structure in C. laticeps using samples from two MPAs and several moderately or severely exploited regions. Despite using analytical tools that are sensitive to detect even subtle genetic structure, we found that this species exists as a single, well-mixed stock throughout its core distribution. The high levels of connectivity identified among sites support the findings of previous studies that have indicated that inshore MPAs are an adequate tool for managing overexploited temperate reef fishes. Even though dispersal of adult C. laticeps out of MPAs is limited, the fact that the large adults in these reserves produce exponentially more offspring than their smaller counterparts in exploited areas makes MPAs a rich source of recruits. We nonetheless caution against concluding that the lack of structure identified in C. laticeps and several other southern African teleosts can be considered to be representative of marine teleosts in this region in general. Many such species are represented in more than one marine biogeographic province and may be comprised of regionally adapted stocks that require individual management.     

Evolution of reproductive features and larval development in the genus Amphiglena (Polychaeta: Sabellidae)

Larval development in Amphiglena mediterranea is direct in the parental tube, and larvae crawl away as young juveniles with a radiolar crown capable of feeding. The non-feeding larvae of A. nathae are initially brooded in the parental tube, and then have a swimming phase before settling and developing a radiolar crown. Developmental data and reproductive features such as sperm structure are combined with other morphological data for an assessment of the cladistic relationships of the species in the genus Amphiglena. Two species of Laonome are included as part of the ingroup. Six genera, representing a sister group to the Laonome/Amphiglena clade, are used as an outgroup. Two most-parsimonious cladograms were found, and the evolution of reproductive features are discussed. The monophyly of Amphiglena is indicated by features previously identified, as well as two features based on sperm morphology. At present a plesiomorphic larval form cannot be identified for the genus Amphiglena. Received: 16 June 1997 / Accepted: 29 January 1998     

Spawning behaviour,egg and larval development of the myctophid fish Benthosema pterotum

The distribution of eggs and early larval stages of the myctophid fish Benthosema pterotum was studied during three cruises to the Gulf of Oman during January to February 1981 and February 1983. The eggs and yolk sac larvae were identified and described after artificial fertilization on board the Research Vessel Dr. Fridtjof Nansen. The mature fish spawn in the Gulf at 300 to 100 m depths during early night. The eggs and smallest larvae were found at 300 to 200 m depths. The eggs hatch within 12 h at 21°C. Hatching occurs before the slightly bouyant eggs reach the upper 50 m of the water column.     

Reproductive behaviour and larval development of Tanais cavolinii (Crustacea: Tanaidacea)

Mature female stages of Tanais cavolinii Milne Edwards are described according to degree of oostegite development. Sexual dimorphism of the chelipeds of copulatory males is explained functionally. An aberrant marsupium consisting of two closed ovisacs with a small slit for sperm transfer protects the brood from sudden unpredictable changes in salinity. Behavioural traits, courtship, copulation, and parental care are described and compared to those of Heterotanais oerstedi Kröyer. The different habitats selected by the two species are reflected by several adaptations in their morphology and behaviour as tube dwellers. A phenomenon unique among invertebrates is that the female T. cavolinii supplies additional yolk to the larvae just before their release. The production and timing of this food supply is described and its adaptive significance is discussed.     

Underwater observations on the reproductive activity of the demoiselleChromis dispilus (Pisces: Pomacentridae)

Reproductive behaviour of the demoiselleChromis dispilus Griffin was studied along the coast and offshore islands of north-eastern New Zealand. Spawning occurred at intervals over the summer months, December to March. Males displayed nursery behaviour and nest-guarding during the incubation period of the eggs. Juvenile growth was initially very rapid. A comparison of reproductive activities with other species ofChromis show a basic similarity in behaviour, although the demoiselle appears to differ in certain behavioural patterns associated with spawning.     

Microstructure of settlement-marks in the otoliths of tropical reef fishes

The morphology and ultrastructure of the otolith settlement-mark was examined in 44 tropical reef-fish species spanning nine families. A classification scheme based on similar otolith characteristics is presented. Three major categories are identified based on changes in increment width and optical qualities of the settlement-mark. Of the 44 species examined, 39 possessed “abrupt” settlement-marks (Type I) characterised by a rapid decrease in increment width (up to 50% reduction) over settlement. Type I settlement-marks were found in all nine families examined. The 39 species spanned the whole range of possible larval durations (Pomacentrus moluccensis, 15 d ± 0 SE; Naso hexacanthus, 91.2 d ± 2.97 SE). Four of the 44 species possessed “zonal” settlement-marks (Type II), featuring a band of increments that are wider than pre-settlement increments. Species in this category are the labrids Corisaygula, Thalassoma bifasciatum, T. lunare and an unidentified acanthurid (Acanthurus sp. 2). One species of acanthurid (N. brevirostris) possessed a “gradual” settlement-mark (Type III), manifest as a gradual decrease in increment width during the settlement period. A possible fourth type was identified from the literature. Gnatholepis thompsoni and Coryphopterus glaucofraenum possessed a settlement-mark with increment widths that increased post-settlement. Available data suggest a poor relationship between the structure of the settlement-mark and the magnitude of metamorphosis (previously reported as internal and external morphological change). Evidence suggests that the increment profile over early development and the increment transitions associated with the settlement event are taxon-specific and may enable late-larval stage fishes to be identified to species level. Received: 21 May 1997 / Accepted: 3 February 1999     

Gametogenesis and larval development in Sabella spallanzanii (Polychaeta: Sabellidae) from the Mediterranean Sea

Sabella spallanzanii (Gmelin) is one of the best known and widely distributed Mediterranean polychaetes, but available data on its general biology has been inferred mostly from populations recently introduced to Australia. In the present paper, data on gametogenesis and larval development of a Mediterranean population from the Gulf of Taranto (Ionian Sea, Italy) are reported. Histological and electron microscopical analysis of gametogenesis showed that oogenesis, a long process beginning soon after spawning in February, is of extraovarian type. By contrast spermatogenesis is a very fast process lasting from September to December, when the coelomic cavity is completely packed with mature gametes and almost devoid of coelomocytes. During the period of spawning, some experiments of in vitro fertilisation were performed, always without success, while some of the collected females naturally spawned fertilised eggs (100%). Particularly interesting was the discovery of a small percentage of eggs which appeared fertilised but unspawned within the coelomic cavity of female. Investigation of the S. spallanzanii life cycle, from the naturally spawned fertilised eggs to embryonic and juvenile stages, was performed by daily microscopical observations and by SEM. Despite the presence of large eggs giving rise to a lecithotrophic larva, a long pelagic phase was observed, the longest found among Sabellidae. The larval development pattern is also described. Settlement began after about 2 weeks; metamorphosis occurred 10 d after settlement, when mucus was secreted abundantly and an external tube was formed. The long larval pelagic period and the development pattern, suggesting a high potential for dispersal, support both the introduction and invasive behaviour of the species in Australian waters. Received: 28 May 1999 / Accepted: 10 December 1999